Reassessment of the generic limits for Hydnellum and Sarcodon (Thelephorales, Basidiomycota)

er-reviewed open-access journal

MycoKeys 54: 31-47 (2019) 6
doi: 10.3897/mycokeys.54.35386 RESEARCH ARTICLE Oo Mycokeys

http://mycokeys.pensoft.net Launched to accelerate biodiversity research

Reassessment of the generic limits for
Hydnellum and Sarcodon
(Thelephorales, Basidiomycota)

Karl-Henrik Larsson'?, Sten Svantesson*?*, Diana Miscevic?,

Urmas Kéljalg®, Ellen Larsson”?

| Natural History Museum, University of Oslo, PO. Box 1172 Blindern, NO 0318 Oslo, Norway
2 Gothenburg Global Biodiversity Centre, PO. Box 461, SE 405 30 Goteborg, Sweden 3 Department of
Biological and Environmental Sciences, University of Gothenburg, PO. Box 461, SE 405 30 Goteborg, Sweden
4 Royal Botanic Gardens Victoria, Birdwood Ave, Melbourne, Victoria 3004, Australia 5 Vastkuststiftelsen,
Sandihamnsvigen 71, SE 434 94 Vallda, Sweden 6 Institute of Ecology and Earth Sciences, 40 Lai Street,
51005 Tartu, Estonia

Corresponding author: Karl-Henrik Larsson (k.h.larsson@nhm.uio.no)

Academic editor: Maria P Martin | Received 11 April 2019 | Accepted 21 May 2019 | Published 10 June 2019

Citation: Larsson K-H, Svantesson S, Miscevic D, Kéljalg U, Larsson E (2019) Reassessment of the generic limits for
Hydnellum and Sarcodon (Thelephorales, Basidiomycota) MycoKeys 54: 31-47. https://doi.org/10.3897/
mycokeys.54.35386

Abstract

DNA sequences from the nuclear LSU and ITS regions were used for phylogenetic analyses of Thelephorales
with a focus on the stipitate hydnoid genera Hydnellum and Sarcodon. Analyses showed that Hydnellum
and Sarcodon are distinct genera but that the current division, based on basidioma texture, makes Sarcodon
paraphyletic with respect to Hydnellum. In order to make genera monophyletic several species are moved
from Sarcodon to Hydnellum and the following new combinations are made: Hydnellum amygdaliolens,
H. fennicum, H. fuligineoviolaceum, H. fuscoindicum, H. glaucopus, H. joeides, H. lepidum, H. lundellii,
H.. martioflavum, H. scabrosum, H. underwoodii, and H. versipelle. Basidiospore size seems to separate the
genera in most cases. Hydnellum species have basidiospore lengths in the range 4.45-6.95 um while the
corresponding range for Sarcodon is 7.4-9 um. S. quercinofibulatus deviates from this pattern with an

average spore length around 6 um. Neotropical Sarcodon species represent a separate evolutionary lineage.

Keywords
Phylogeny, stipitate hydnoid, taxonomy, Thelephorales, tooth fungi

Copyright Karl-Henrik Larsson et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

32 Karl-Henrik Larsson et al. / MycoKeys 54: 31-47 (2019)

Introduction

The order Thelephorales is a distinctive lineage of Agaricomycetes, well-known for
its almost ubiquitous ectomycorrhizal life style (Tedersoo et al. 2010). Several species
have stipitate hydnoid basidiomata (Fig. 1). They have traditionally been divided into
four genera, Phellodon and Bankera with hyaline basidiospores, and Hydnellum and
Sarcodon with yellow to brown tinted basidiospores (Maas Geesteranus 1975). In
both cases the genera within each pair differ in basidioma structure, with Phellodon
and Hydnellum being hard and dry, and Bankera and Sarcodon forming softer, fleshier
basidiomata. This difference in texture is, however, difficult to assess and a series of
recent molecular phylogenetic analyses, as outlined below, have indicated that the
traditional, morphology-based generic limits are equivocal.

In a recent comprehensive study of stipitate hydnoid species from south-eastern North
America, Baird et al. (2013) found that Bankera could not be separated from Phellodon
and the genera were hence combined into a more comprehensive Phellodon. The same
study suggested that the generic limits of Sarcodon and Hydnellum need reassessment.

Nitare and Hégberg (2012) examined the Nordic species of Sarcodon and included
a preliminary molecular phylogeny for the species accepted in Sarcodon. Hydnellum
species were also included in non-published test runs and found to be nested among
Sarcodon species. They concluded that revisions of limits of both genera were probably
necessary. Miscevic (2013) expanded on the results in Nitare and Hégberg (2012) by
including more sequences for each species and by including a selection of Hydnellum
species in published phylogenies. The results were in congruence with Baird et al.
(2013) with regard to overall tree topology and again the conclusion was that the limits
of Sarcodon and Hydnellum need further study. A recent phylogenetic overview of
Thelephorales (Vizzini et al. 2016) and a study of Hydnellum from the Mediterranean
region (Loizides et al. 2016) came to similar conclusions, although Vizzini et al. (2016)
did not include sequences from several Neotropical Sarcodon species described by
Grupe et al. (2015, 2016).

In this paper we analyse ITS and nuclear LSU sequences from a wide selection of
Thelephorales species with a focus on Hydnellum and Sarcodon in order to resolve the
relationship between these two genera. We also make some nomenclatural changes that
follow from the revision of genus circumscriptions. We demonstrate that Neotropical
Sarcodon species do not cluster with temperate and boreal species and may be warranted
as one or more new genera with more data.

Methods

For the phylogenetic analyses we compiled two datasets. ‘The first dataset consists of
nuclear LSU sequences from most genera in Thelephorales and from a majority of
the Hydnellum and Sarcodon species occurring in Europe. For our two target genera
we chose only sequences generated for this study from recently collected basidiomata.

Reassessment of the generic limits for Hydnellum and Sarcodon 33

Figure |. Fruiting bodies of Hydnellum and Sarcodon A Hydnellum suaveolens B H. aurantiacum

C H. ferrugineum D Sarcodon imbricatus.

We deliberately excluded sequences from specimens identified as H. concrescens or
H. scrobiculatum since these names seem to cover more than just two species and it
is currently unclear how the names should be applied (Ainsworth et al. 2010). Since
this study is positioned as a revision of the genus limits we were more interested in
sequence quality control than a complete coverage of all species reported from Europe.

For our second dataset we chose a different strategy. Here we included ITS sequences
from all Hydnellum and Sarcodon species represented among our own sequences and in
GenBank as of December 1, 2018. The reason is that many species, and especially the
recently described species from tropical regions, are only available as ITS sequences.
However, we made no attempt to verify the identifications given in GenBank and do
not endorse them as correct.

DNA was extracted from recent dried collections of basidiomata from North Eu-
rope. Voucher numbers, herbarium location, and GenBank numbers are given in Table
1. DNA extraction and PCR protocols follow Larsson et al (2018). Sequencing was ei-
ther done in-house at University of Oslo, or as a commercial service by Macrogen Inc.,
South Korea. Assembly of chromatograms was done with Sequencher 5.2.4 (Gene
Codes Co., Ann Arbor). Aligning was performed either manually using the editor in
PAUP* 4.0a (Swofford 2002) or the software ALIVIEW 1.18 (Larsson 2014), or au-
tomatically utilising the L-INS-i strategy as implemented in MAFFT v. 7.017 (Katoh
and Standley 2013), followed by manual adjustment.

34 Karl-Henrik Larsson et al. / MycoKeys 54: 31-47 (2019)

Table I. Specimens sequenced or downloaded from GenBank. Herbarium acronyms follow Thiers.

Sequences generated for this study are marked in bold.

Species

Amaurodon aquicoeruleus Agerer

Amaurodon viridis (Alb. & Schwein.:Fr.) J.Schrot
Bankera fuligineoalba (J.C.Schmidt:Fr.) Pouzar
Bankera violascens (Alb. & Schwein.:Fr.) Pouzar
Boletopsis leucomelaena (Pers.:Fr.) Fayod
Hydnellum aurantiacum (Batsch:Fr.) P.Karst.
Hydnellum aurantiacum

Hydnellum aurantiacum

Hydnellum auratile (Britzelm.) Maas Geest.
Hydnellum auratile

Hydnellum auratile

Hydnellum caeruleum (Hornem.:Fr.) P.Karst.
Hydnellum caeruleum

Hydnellum caeruleum

Hydnellum complicatum Banker

Hydnellum concrescens (Pers.) Banker
Hydnellum cristatum (G.EAtk.) Stalpers
Hydnellum cumulatum K.A.Harrison
Hydnellum cyanopodium K.A.Harrison
Hydnellum diabolus Banker

Hydnellum dianthifolium Loizides, Arnolds & P.-A.Moreau
Hydnellum earlianum Banker

Hydnellum ferrugineum (Fr.:Fr.) P.Karst.
Hydnellum ferrugineum

Hydnellum ferrugineum

Hydnellum ferrugipes Coker

Hydnellum geogenium (Fr.) Banker

Hydnellum geogenium

Hydnellum geogenium

Hydnellum gracilipes (P.Karst.) P.Karst.
Hydnellum gracilipes

Hydnellum mirabile (Fr.) P.Karst.

Hydnellum mirabile

Hydnellum mirabile

Hydnellum peckii Banker

Hydnellum peckii

Hydnellum peckii

Hydnellum pineticola K.A.Harrison

Hydnellum piperatum Maas Geest.

Hydnellum regium K.A.Harrison

Hydnellum scleropodium K.A.Harrison
Hydnellum scrobiculatum (Fr.) P.Karst.
Hydnellum spongiosipes (Peck) Pouzar
Hydnellum suaveolens (Scop.:Fr.) P.Karst.
Hydnellum suaveolens

Hydnellum suaveolens

Hydnellum subsuccosum K.A.Harrison
Lenzitopsis daii L.W.Zhou & Kéljalg
Lenzitopsis oxycedri Malengon & Bertault
Odontia fibrosa (Berk. & M.A.Curtis) Kéljalg
Phellodon cf niger

Phellodon tomentosus (L.:Fr.) Banker
Pseudotomentella flavovirens (Hohn. & Litsch.) Svréek
Sarcodon amygdaliolens Rubio Casas, Rubio Roldan & Catala
Sarcodon aspratus (Berk.) S.Ito

Voucher

Agerer & Bougher
KH Larsson 14947b
E Larsson 400-13
MV 130902
M Krikorev 140912
RG Carlsson 08-105
E Bendiksen 177-07
O-F-295029
O-F-294095
O-F-242763
J Nitare 110926
O-F-291490
E Bendiksen 575-11
E Bendiksen 584-11
REB 71
K(M) 134463
REB 169
SE Westmoreland 69
SE Westmoreland 85
KAH 13873
ML61211HY
REB 375
O-F-297319
E Larsson 356-16
E Larsson 197-14
REB 176
O-F-66379
O-F-296213
E Bendiksen 526-11
E Larsson 219-11
GB-0113779
RG Carlsson 11-119
E Larsson 170-14
S Lund 140912
S Svantesson 328
E Larsson 174-14
E Bendiksen 567-11
RB 94
REB 322
SE Westmoreland 93
REB 3
REB 78
REB 52
E Larsson 139-09
E Larsson 8-14
S Svantesson 877
REB 10
Yuan 2959
KH Larsson 15304
TU115028
E Larsson 35-14
E Bendiksen 118-10
KH Larsson 16190
SC 2011

Herb.

GenBank number

ITS LSU
AM490944 AM490944
MKG602707 MKG602707
MK602708 MK602708
MKG602709 MK602709
MK602710 MK602710
MK602711 MK602711
MK602712 MK602712
MKG602713 MK602713
MK602714 MK602714
MK602715 MK602715
MK602716 MK602716
MKG602717 MKG602717
MKG602718 MK602718
MKG602719 MKG602719
KC571711
EU784267
JN135174
AY569026
AY569027
AF351863
KX619419
JN135179
MK602720 MK602720
MK602721 MK602721
MK602722 MK602722
KC571727
MK602723 MK602723
MK602724 MK602724
MK602725 MK602725
MK602726 MK602726
MK602727 MK602727
MK602728 MK602728
MK602729 MK602729
MK602730 MK602730
MK602731 MK602731
MK602732 MK602732
MK602733 MK602733
KC571734
JN135173
AY569031
JN135186
JN135181
JN135184
MK602734 MK602734
MK602735 MK602735
MK602736 MK602736
JN135178
JN169799 JN169793
MK602774 MK602774
MK602775 MK602775
MK602782 MK602782
MK602781 MK602781
MK602780 MK602780
JN376763
DQ448877

Reassessment of the generic limits for Hydnellum and Sarcodon

Species

Sarcodon atroviridis (Morgan) Banker
Sarcodon atroviridis

Sarcodon bairdii A.C.Grupe & Vasco-Pal.
Sarcodon colombiensis A.C.Grupe & Vasco-Pal.
Sarcodon fennicus (P.Karst.) P.Karst.

Sarcodon fennicus

Sarcodon fennicus

Sarcodon fuligineoviolaceus (Kalchbr.) Pat.
Sarcodon fuligineoviolaceus

Sarcodon fuligineoviolaceus

Sarcodon fuscoindicus (K.A.Harrison) Maas Geest.
Sarcodon glaucopus Maas Geest. & Nannf.
Sarcodon glaucopus

Sarcodon glaucopus

Sarcodon imbricatus (L.:Fr.) P.Karst.

Sarcodon imbricatus

Sarcodon imbricatus

Sarcodon joeides (Pass.) Bataille

Sarcodon joeides

Sarcodon joeides

Sarcodon joeides

Sarcodon lepidus Maas Geest.

Sarcodon lepidus

Sarcodon lepidus

Sarcodon leucopus (Pers.) Maas Geest. & Nannf.
Sarcodon leucopus

Sarcodon leucopus

Sarcodon lundellii Maas Geest. & Nannf.
Sarcodon lundellii

Sarcodon lundellii

Sarcodon martioflavus (Snell, K.A.Harrison & H.A.C.Jacks.)

Maas Geest.

Sarcodon martioflavus

Sarcodon martioflavus

Sarcodon pakaraimensis A.C.Grupe & T.W.Henkel
Sarcodon pallidogriseus A.C.Grupe & Vasco-Pal.
Sarcodon portoricensis A.C.Grupe & T.J.Baroni
Sarcodon quercophilus A.C.Grupe & Lodge

Sarcodon quercinofibulatus Pérez-De-Greg., Macau & J.Carbé

Sarcodon rufobrunneus A.C.Grupe & Vasco-Pal.
Sarcodon scabripes (Peck.) Banker

Sarcodon scabrosus (Fr.) P.Karst.

Sarcodon scabrosus

Sarcodon scabrosus

Sarcodon squamosus (Schaeff.) Quél.

Sarcodon squamosus

Sarcodon squamosus

Sarcodon umbilicatus A.C.Grupe, T.J.Baroni & Lodge

Sarcodon underwoodii Banker

Sarcodon versipellis (Fr.) Nikol.

Sarcodon versipellis

Sarcodon versipellis

Sistotrema brinkmannii (Bres.) J.Erikss.
Steccherinum ochraceum (J.E.Gmel.:Fr.) Gray
Thelephora caryophyllea (Schaeff.:Fr.) Pers.
Thelephora terrestris Ehrh.:Fr.

Tomentella stuposa (Link) Stalpers
Tomentellopsis pulchella Kéljalg & Bernicchia

Voucher

REB 104
REB 61
Vasco 990
Vasco 2084
S Westerberg 110909
O-F-242833
O-F-204087
LA 120818
B Nylén 130918
A Molia 160-2011
OSC 113622
RG Carlsson 13-060
J Nitare 060916
A Edvinson 110926
S Svantesson 355
J Rova 140829-2
E Larsson 384-10
RG Carlsson 11-090
K Hjortstam 17589
J Nitare 110829
REB 270
E Grundel 110916
RG Carlsson 10-065
J Nitare 110829
O-F-296944
O-F-296099
P Hedberg 080811
L&A Stridvall 06-049
O-F-242639
O-F-295814
A Delin 110804

O-F-242435
O-F-242872
T Henkel 9554
Vasco 989
TG Baroni 8776
CFMR-BZ-3833
JC 20090718-2
Vasco 1989
REB 351
O-F-295824
O-F-292320
O-F-360777
O-F-177452
E Larsson 248-12
O-F-295554
TJ Baroni 10201
REB 50
RG Carlsson 13-057
RG Carlsson 11-085
E Bendiksen 164-07
KH Larsson 14078
KH Larsson 11902
E Larsson 89-09S
E Larsson 295-13
Th-0764
KH Larsson 16366

Herb.

TENN

GenBank number
ITS LSU

JN135190

KC571768

KR698938

KP972654
MK602739 MK602739
MK602738 MK602738
MK602737 MK602737
MK602740 MK602740
MK602741 MK602741
MK602742 MK602742
EU669228
MK602743 MK602743
MK602744 MK602744
MK602745 MK602745
MK602748 MK602748
MKG602746 MKG602746
MKG602747 MK602747
MKG602749 MKG602749
MKG602750 MK602750
MK602751 MK602751
KC571772
MK602753 MK602753
MK602752 MK602752
MK602754 MK602754
MKG02756 MKG602756
MK602755 MK602755
MK602757 MKG602757
MK602758 MK602758
MK602759 MK602759
MK602760 MK602760
MKG602763 MK602763
MK602762 MK602762
MK602761 MK602761
KM668103

KR698939

KM668100

KM668101

JX271818 MK602773
KR698937

JN135191
MK602764 MK602764
MKG02766 MKG02766
MKG602765 MKG602765
MK602768 MK602768
MKG602767 MKG602767
MKG602769 MKG602769
KM668102

KC571781
MK602771 MK602771
MK602772 MK602772
MKG602770 MKG602770
KF218967 KF218967
JQ031130 JQ031130
MKG602776 MK602776
MK602777 MK602777
MK602778 MK602778
MK602779 MK602779

35

36 Karl-Henrik Larsson et al. / MycoKeys 54: 31-47 (2019)

In the phylogenetic analyses we assumed the following minimal partitions
for the nrDNA region: ITS1, 5.88, ITS2 and LSU (approximately 1200 bases of
the 5’ end). Two datasets were analysed separately: an LSU dataset only including
the LSU region, and an ITS dataset including ITS1, 5.8S and ITS2. We used the
automated best-fit tests implemented in PAUP* 4.0a (Swofford 2002) to select
optimal substitution models for each complete, non-partitioned dataset (PHYML)
and optimal substitution model partitions for each minimal partition (BEAST).
Models and partitions were chosen based on BIC score for the BEAST analysis
and AICc score for the PHYML analysis. All tests were conducted using three
substitution schemes and evaluated substitution models with equal and gamma-
distributed among-site rate variation. The tests for the PHYML analysis also
evaluated substitution models with invariant sites. The following partitions and
models had the highest ranking, according to BIC: ITS1+ITS2 (GTR+G), 5.88
(K80+G), LSU (GTR+G). According to AICc the GT R+I+G model provided the
best fit for both the ITS and the LSU datasets.

To generate Bayesian phylogenetic trees (BI) from the alignments we used BEAST
2.4.7 (Bouckaert et al. 2014). We prepared the xml-files for the BEAST 2 runs in
BEAUTI 2.4.7 (Bouckaert et al. 2014). We set the substitution model to GITR+G for
the LSU run. In the ITS run we set it to HKY+G for 5.8S, since it is the most similar
model to K80+G available in the program. Test runs revealed convergence problems
due to insufficient data for some substitution rates in the GIR+G model initially used
for the ITS1+ITS2 partition, and it was hence changed to HKY+G. In the ITS run the
substitution rate of both partitions were estimated independently. We set the trees of
the minimal nrDNA partitions as linked in this analysis and the clock models as un-
linked. A lognormal, relaxed clock model was assumed for each partition, as test runs
had shown that all partitions had a coefficient of variation well above 0.1 (i.e. implying
a relatively high rate variation among branches). ‘The clock rate of each partition was
estimated in the runs, using a lognormal prior with a mean set to one in real space.
We set the growth rate prior to lognormal, with a mean of 5 and a standard deviation
of 2. We ran the Markov Chain Monte Carlo (MCMC) chains of both datasets for
20 million generations with tree and parameter files sampled every 1,000 generations.
The analyses all converged well in advance of the 10 % burn-in threshold, had ESS
values well above 200 for all parameters, and chain mixing was found to be satisfac-
tory as assessed in TRACER 1.6.0 (Rambaut et al. 2014). After discarding the burn-in
trees, maximum clade credibility trees were identified by TREEANNOTATOR 2.4.7
(Bouckaert et al. 2014).

To generate Maximum Likelihood (ML) gene trees we used PHYML 3.1 (Guin-
don et al. 2010). We set the substitution model to GT R+I+G for both the ITS and
LSU datasets. Tree topology search was conducted using NNI+SPR, with ten random
starting trees. Non-parametric bootstrap analyses with 1000 replicates were performed
on the resulting trees.

Reassessment of the generic limits for Hydnellum and Sarcodon oF

Results

Seventy-five Thelephorales specimens from the genera Amaurodon, Bankera, Boletopsis,
Hydnellum, Lenzitopsis, Phellodon, Pseudotomentella, Sarcodon, Thelephora, Tomentella,
and Yomentellopsis, were sequenced for this study. In addition, 39 sequences were
downloaded from public databases (GenBank, UNITE) including outgroup sequences
of Steccherinum ochraceum (Polyporales) and Sistotrema brinkmannii (Cantharellales)
included in the LSU dataset. The ITS analyses were rooted by the default method
(BEAST) or left unrooted (PHYML).

The aligned LSU dataset consisted of 1443 nucleotide positions. After exclusion of
ambiguous regions 1377 positions remained for the analyses. BI returned a tree where the
focus genera Hydnellum and Sarcodon are distributed over two strongly supported clades.
The larger of these clades includes the type of Hydnellum, H. suaveolens, and an additional
17 species, all except one forming strongly supported terminal clades. Nine of these taxa are
currently placed in Sarcodon. With a few exceptions the relationships within Hydnellum are
not resolved. H. aurantiacum and H. auratile are recovered as a strongly supported group;
Sarcodon scabrosus and S. fennicus are grouped with 0.97 posterior probability support; S.

fuligineoviolaceus, S. glaucopus, and S. joeides form a subclade with 0.97 posterior probability

support; and finally H. suaveolens and S. versipellis form a strongly supported clade. The
type of Sarcodon, S. imbricatus, and three other species form the second main clade. The
three sequences of S. imbricatus cluster together but the clade is unsupported. Hydnellum
and Sarcodon are recovered as sister clades but the support for this arrangement is weak.

For target taxa the ML tree is essentially similar to the BI tree with strong support
for the similarly composed Hydnellum and Sarcodon clades (Fig. 2). As for the BI
analysis the relationships among species within Hydnellum and Sarcodon are not
resolved except for a weak to moderate support for grouping H. aurantiacum with H.
auratile and H. suaveolens with S. versipellis. S. fuligineoviolaceus, S. glaucopus, and S.

joeides also group together in the ML tree but without support. Again S. imbricatus
does not get support and is not separated from S. quercinofibulatus.

The aligned ITS dataset consisted of 1068 nucleotide positions of which 505
remained for the analyses after removal of ambiguous regions. Bayesian inference
produced a tree with two strongly supported clades (Fig. 3). The smaller one, which
we here informally call “Neosarcodon”, contains nine Sarcodon species, all with a
distribution in the tropical and subtropical Americas. Remaining Hydnellum and
Sarcodon taxa, including both type species, formed the other clade. Within the latter
clade two subclades are visible, corresponding to the genera Hydnellum and Sarcodon,
and with the same delimitation as in the LSU trees. Only the Sarcodon subclade has
strong support. Within each larger clade several groups of taxa received moderate to
strong support. The reader is referred to Fig. 2 for further details.

The ML tree recovered the same two main clades with strong support but could
not resolve the relationships within the larger Hydnellum/Sarcodon clade. In the ML

38 Karl-Henrik Larsson et al. / MycoKeys 54: 31-47 (2019)

Hydnellum aurantiacum MK602711
0.99/87 Hydnellum aurantiacum MK602712
87 Hydnellum aurantiacum MK602713
Hydnellum auratile MK602713
Hydnellum auratile MK602714
Hydnellum auratile MK602715
99 Sarcodon scabrosus MK602764
Sarcodon scabrosus MK602766
Sarcodon scabrosus MK602765
0.97/- Sarcodon fennicus MK602739
‘ Sarcodon fennicus MK602738
Sarcodon fennicus MK602737
Sarcodon lundellii MK602758
Sarcodon lundellii MK602759
Sarcodon lundellii MK602760
86 Sarcodon glaucopus MK602743
Sarcodon glaucopus MK602744
Sarcodon glaucopus MK602745
Sarcodon fuligineoviolaceus MK602740
0.97/- Sarcodon fuligineoviolaceaus MK602741
r 84 Sarcodon fuligineoviolaceus MK602742
Sarcodon joeides MK602749
Sarcodon joeides MK602750
91! Sarcodon joeides MK602751
Hydnellum mirabile MK602728
Hydnellum mirabile MK602729
Hydnellum mirabile MK602730
Sarcodon lepidus MK602753
Sarcodon lepidus MK602752
Sarcodon lepidus MK602754
Hydnellum gracilipes MK602726
Hydnellum gracilipes MK602727
Sarcodon martioflavus MK602763
Sarcodon martioflavus MK602762
Sarcodon martioflavus MK602761
Hydnellum caeruleum MK602717
Hydnellum caeruleum MK602718
Hydnellum caeruleum MK602719
Hydnellum geogenium MK602723
Hydnellum geogenium MK602724
Hydnellum geogenium MK602725
90 Hydnellum peckii MK602731
97 Hydnellum peckii MK602732
Hydnellum peckii MK602733
Hydnellum suaveolens MK602734
Hydnellum suaveolens MK602735
74 Hydnellum suaveolens MK602736
Sarcodon versipellis MK602771
Sarcodon versipellis MK602772
Sarcodon versipellis MK602770
0.93/- Hydnellum ferrugineum MK602720
. Hydnellum ferrugineum MK602721
Hydnellum ferrugineum MK602722
99 Sarcodon leucopus MK602755
Sarcodon leucopus MK602756
Sarcodon leucopus MK602757
Sarcodon quercinofibulatus MK602773
Sarcodon imbricatus MK602748
97 Sarcodon imbricatus MK602746
Sarcodon imbricatus MK602747
Sarcodon squamosus MK602768
Sarcodon squamosus MK602767
97' Sarcodon squamosus MK602769
Boletopsis leucomelaena MK602710
98 Lenzitopsis daii JN169799/JN169793
Lenzitopsis oxycedri MK602774
99 -/7) Thelephora caryophyllea MK602776
75 Thelephora terrestris MK602777
Tomentella stuposa MK602778
Odontia fibrosa MK602775
0.98/96 Pseudotomentella flavovirens MK602780
: 77,— Bankera fuligineoalba MK602708
98 Bankera violescens MK602709
Phellodon cf niger MK602782
-/82 Phellodon tomentosus MK602781
Tomentellopsis pulchella MK602779
Amaurodon viridis MK602707
Amaurodon aquicoeruleus AM490944
Steccherinum ochraceum JQ031130

Sistotrema brinkmannii KF218967

Hydnellum

Sarcodon

0.1

Figure 2. Maximum likelihood analyses of LSU dataset for Thelephorales. Branches in bold have a
posterior probability value of 1 in Bayesian inference and 100% bootstrap support in ML analysis, if
not otherwise indicated by a figure. Lower support values on other branches are indicated by figures.

Steccherinum ochraceum and Sistotrema brinkmannii are used as outgroup (branch lengths shortened).

Reassessment of the generic limits for Hydnellum and Sarcodon

-/70

1/99
0.99/-

1/76
1/99

0.98/-

1/98

1/96

0.99/78

1/94
1/100

0.97/-

0.96/61

0.98/74 =

0.92/na
0.97

1/na
/52

1/87
1/98

1/69 0.98/81
0.94/61

1/100
1/98

0.05 length units

Sarcodon fuligineoviolaceus MK602740
Sarcodon fuscoindicus EU669228
Sarcodon joeides KC571772
Sarcodon joeides MK602751
Sarcodon glaucopus MK602745
Hydnellum mirabile MK602728
Hydnellum peckii MK602733
Hydnellum regium AY569031
Hydnellum gracilipes MK602727
Hydnellum piperatum JN135173
Hydnellum cristatum JN1351 74
Sarcodon lundellii MK602760
Sarcodon versipellis MK602771
Hydnellum suaveolens MK602735
Hydnellum cumulatum AY569026
Hydnellum ferrugineum MK602721
Hydnellum spongiosipes JN135184
Hydnellum pineticola KC571734
Hydnellum diabolus AF351863
Hydnellum geogenium MK602723
Hydnellum cyanopodium AY569027
Hydnellum scleropodium JN135186
Hydnellum caeruleum MK602719
Hydnellum ferrugipes KC571727
Sarcodon martioflavus MK602762
Hydnellum concrescens EU/84267
Hydnellum subsuccosum JN135178
Hydnellum scrobiculatum JN135181
Hydnellum dianthifolium KX619419
Sarcodon lepidus MK602753
Sarcodon fennicus MK602737
Sarcodon scabrosus MK602766
Sarcodon amygdaliolens JN376763
Sarcodon underwoodii KC571781
Hydnellum aurantiacum MK602711
Hydnellum earlianum JN135179
Hydnellum auratile MK602714
Hydnellum complicatum KC571711
Sarcodon imbricatus MK602747
Sarcodon squamosus MK602767
Sarcodon quercinofibulatus JX271818
Sarcodon aspratus DQ448877
Sarcodon scabripes JN135191
Sarcodon leucopus MK602755
Sarcodon atroviridis JN135190
Sarcodon atroviridis KC571768
Sarcodon portoricensis KM668100
Sarcodon quercophilus KM668101
Sarcodon umbilicatus KM668102
Sarcodon rufobrunneus KR698937
Sarcodon pakaraimensis KM668103
Sarcodon columbiensis KP972654
Sarcodon bairdii KR698938
Sarcodon pallidogriseus KR698939

59

Hydnellum

Sarcodon

“Neosarcodon”

Figure 3. Ultrametric default rooted BEAST tree of ITS dataset for Hydnellum and Sarcodon. Posterior

probability values and bootstrap percent support from ML analysis are indicated by figures; na = not applicable.

40 Karl-Henrik Larsson et al. / MycoKeys 54: 31-47 (2019)

tree the clade corresponding to Hydnellum in the LSU tree is correctly identified but
not supported while the clade corresponding to Sarcodon appears polyphyletic.

Based on these results we hereby revise the limits of the two genera by moving a
number of species from Sarcodon to Hydnellum. Consequently the genus description
for Hydnellum must be emended while the genus description for Sarcodon can
remain unaltered.

Taxonomy

Hydnellum P.Karst., Meddn Soc. Fauna Flora fenn. 5: 41 (1879).

Type species. Hydnellum suaveolens (Scop.:Fr.) P.Karst. (1879)

Basionym. Hydnum suaveolens Scop.:Fr. (1772)

Basidiomata with pileus and stipe, single or concrescent; pileus thin to thick, at
first smooth and velutinous, when mature felted, fibrillose, scaly, ridged, or irregularly
pitted and scrupose, mostly brownish but also with white, olive yellowish, orange, pur-
plish or bluish colours, often concentrically zonate; stipe narrow to thick, solid, mostly
short; hymenophore hydnoid, usually strongly decurrent; context from soft and brittle
to corky or woody; hyphal system monomitic, septa with or without clamps, context
hyphae inflated or not; cystidia lacking; basidia narrowly clavate, producing four ster-
igmata; basidiospores with irregular outline, more or less lobed, verrucose, brownish.
Terrestrial, forming ectomycorrhiza with forest trees.

Hydnellum amygdaliolens (Rubio Casas, Rubio Roldan & Catala) E.Larss.,
K.H.Larss. & Kéljalg, comb. nov.
MycoBank No.: MB830570

Basionym. Sarcodon amygdaliolens Rubio Casas, Rubio Roldan & Catala, Boln Soc.
Micol. Madrid 35: 44-45. 2011. Holotype: Spain, Tamajén, Barranco la Jara. L.
Rubio-Casas & L. Rubio-Roldan, AH 42113.

Hydnellum fennicum (P.Karst.) E.Larss., K.H.Larss. & Koljalg, comb. nov.
MycoBank No.: MB830571

Basionym. Sarcodon scabrosus var. fennicus P.Karst., Bidr. Kann. Finl. Nat. Folk 37:
104. 1882. Type: not indicated (neotype: H, designated by Maas Geesteranus &
Nannfeldt 1969: 406)

Reassessment of the generic limits for Hydnellum and Sarcodon 41

Hydnellum fuligineoviolaceum (Kalchbr.) E.Larss., K.H.Larss. & K6ljalg, comb. nov.
MycoBank No.: MB830572

Basionym. Hydnum fuligineoviolaceum Kalchbr., in Fries, Hymenomyc. eur. (Upsaliae):
602. 1874. Holotype: Slovakia, Presovsky kraj, Olaszi. C. Kalchbrenner, UPS F-173546.

Hydnellum fuscoindicum (K.A.Harrison) E.Larss., K.H.Larss. & Koljalg, comb. nov.
MycoBank No.: MB830573

Basionym. Hydnum fuscoindicum K.A.Harrison, Can. J. Bot. 42: 1213. 1964.
Holotype: USA, Washington, Olympic Nat. Park, A.-H. Smith. MICH 10847.

Hydnellum glaucopus (Maas Geest. & Nannf.) E.Larss., K.H.Larss. & Koljalg,
comb. nov.

MycoBank No.: MB830574

Basionym. Sarcodon glaucopus Maas Geest. & Nannf., Svensk bot. Tidskr. 63: 407.
1969. Holotype: Sweden, Uppland, Borje par., J. Eriksson. UPS F-013955.

Hydnellum joeides (Pass.) E.Larss., K.H.Larss. & Koljalg, comb. nov.
MycoBank No.: MB830575

Basionym. Hydnum joeides Pass., Nuovo G. bot. ital. 4: 157. 1872. Holotype: Italy,
Emilia-Romagna, Collecchio, G. Passerini. PAD.

Hydnellum lepidum (Maas Geest.) E. Larss., K.H.Larss. & Kéljalg, comb. nov.
MycoBank No.: MB830576

Basionym. Sarcodon lepidus Maas Geest., Verh. K. ned. Akad. Wet., tweede sect. 65:
105. 1975. Holotype: The Netherlands, Lochem, Ampsen, G. & H. Piepenbroek. L.

Hydnellum lundellit (Maas Geest. & Nannf.) E.Larss., K.H.Larss. & Koljalg,
comb. nov.

MycoBank No.: MB830577

Basionym. Sarcodon lundellii Maas Geest. & Nannf., Svensk bot. Tidskr. 63: 421.
1969. Type: Sweden, Uppland, Storvreta, S. Lundell & J.A. Nannfeldt, distributed

42 Karl-Henrik Larsson et al. | MycoKeys 54: 31-47 (2019)

in S. Lundell & J.A. Nannfeldt Fungi exs. suec. as number 252 (lectotype, designated
here, UPS F-010975; MycoBank No.: MBT387081). The UPS herbarium has two
copies of the exsiccate and the specimens of H. /undellii are registered as F-010975 and
F-013956, respectively. From F-010975 an ITS2 sequence has been generated [Gen-
Bank MK753037] and this specimen is here selected as lectotype).

Hydnellum martioflavum (Snell, K.A.Harrison & H.A.C.Jacks.) E.Larss.,
K.H.Larss. & Kéljalg, comb. nov.
MycoBank No.: MB830578

Basionym. Hydnum martioflavum Snell, K.A.Harrison & H.A.C.Jacks., Lloydia 25:
161. 1962. Holotype: Canada, Quebec, Ste Anne de la Pocatiére, H.A.C. Jackson &
W.H. Snell 13 Sep. 1954, BPI 259438.

Hydnellum scabrosum (Ft.) E.Larss., K.H.Larss. & Koljalg, comb. nov.
MycoBank No.: MB830579

Basionym. Hydnum scabrosum Fr., Anteckn. Sver. Atl. Svamp.: 62. 1836. Type: not
indicated (neotype: Sweden, Smaland, Femsjé, S. Lundell, UPS F-013954, designated
by Maas Geesteranus & Nannfeldt 1969: 426)

Hydnellum underwoodii (Banker) E.Larss., K.H.Larss. & Koljalg, comb. nov.
MycoBank No.: MB830580

Basionym. Sarcodon underwoodii Banker, Mem. Torrey bot. Club 12: 147. 1906.
Holotype: USA, Connecticut, NY 776131.

Hydnellum versipelle (Fr.) E.Larss., K.H.Larss. & Koéljalg, comb. nov.
MycoBank No.: MB830581

Basionym. Hydnum versipelle Fr., Ofvers. K. Svensk. Vetensk.-Akad. Férhandl. 18(1):
31. 1861. Type: not indicated (neotype: Sweden, Uppland, Danmark par., J. Eriksson &
H. Nilsson, UPS F-013958, designated by Maas Geesteranus & Nannfeldt 1969: 430)

Sarcodon Quél. ex P.Karst., Revue mycol., Toulouse 3 (no. 9): 20 (1881).

Type species. Sarcodon imbricatus (L.:Fr.) P.Karst. (1881)
Basionym. Hydnum imbricatum L.:Fr. (1753).

Reassessment of the generic limits for Hydnellum and Sarcodon 43

Basidiomata with pileus and stipe, single or concrescent; pileus thin to thick, at
first smooth and velutinous, when mature smooth or scaly, brownish; stipe thick,
solid, mostly short; hymenophore hydnoid, usually strongly decurrent; context soft
and brittle; hyphal system monomitic, septa with clamps, context hyphae inflated;
cystidia lacking; basidia narrowly clavate, producing four sterigmata; basidiospores
with irregular outline, more or less lobed, verrucose, brownish. Terrestrial, forming
ectomycorrhiza with forest trees.

Discussion

In this paper we show that the current morphology-based concepts of Sarcodon and
Hydnellum do not correspond to monophyletic subgroups within the Thelephorales.
The characters traditionally used to separate the two genera do not reflect true
relationships. These characters, however, are vague and open to subjectivity; hence it
is not surprising that they have now been shown to be unreliable. Maas Geesteranus
(1975) pointed to the context structure and consistency as the main differentiating
character. For Hydnellum he describes the context as “... fibrillose, soft or tough, corky
to woody, more or less duplex, zoned, ...” and hyphae are said to be “...usually not
inflating ...”. In Sarcodon the same structures are described as “... fleshy, brittle, soft or
firm (never corky or woody), not duplex, not zoned ...” and “...hyphae inflating ...”.
While these morphological characteristics remain true for Sarcodon, the corresponding
descriptions for Hydnellum had to be emended.

Instead of context structure it seems that average basidiospore size may in most cases
offer a possibility to separate a Sarcodon species from one belonging to Hydnellum. Table
2 summarizes basidiospore measurements from the literature. Average basidiospore
lengths in Hydnellum fall between 4.45 and 6.95 um while the same figures for Sarcodon
are 7.4 and 9 um, ornamentation excluded. However, S. quercinofibulatus clearly deviates
from this pattern. According to measurements in the protologue (Pérez-de-Gregorio et
al. 2011) and in Vizzini et al. (2013) average basidiospore length was measured to 6.95
and 7.0, respectively, but then included the ornamentation. Measurements excluding
ornamentation would be approximately 1 pm less. Clearly, for S. guercinofibulatus
basidiospore length alone will not be decisive for genus placement.

Not all sequences from species described as Sarcodon spp. were recovered within
either Sarcodon or Hydnellum. In our ITS-only analyses nine species formed a well-
supported clade of their own, separated from Sarcodon sensu stricto and Hydnellum
(Fig. 3). This clade, here informally called “Neosarcodon’”, contains species collected in
tropical and subtropical regions of the Western Hemisphere and may represent one or
several distinct genera. However, further analyses based on an expanded dataset using
more conservative molecular markers would be required to definitely identify any new
higher taxa in the group.

The failure to generate support for Sarcodon and Hydnellum in the ITS-only
analyses reflects the large genetical distances present among the species within this

44 Karl-Henrik Larsson et al. | MycoKeys 54: 31-47 (2019)

Table 2. Basidiospore measurements for Hydnellum and Sarcodon from the literature. Sources: B = Baird
et al. (2013), M = Maas Geesteranus (1975), J = Johannesson et al. (1999). All measurements exclude
ornamentation. For species treated in this paper names follow our new classification. For other species

names are according to cited authors.

Species Measurements Mean length
Hydnellum aurantiacum (M) (5.8-)6-6.7 x (4-)4.3-4.9 635
Hydnellum auratile (M) 4.9-5.8 x 3.6-4.5 5235
Hydnellum caeruleum (M) 5.4-6(-6.3) x 3.4-4.3 5.70
Hydnellum compactum (Pers.:Fr.) P. Karst. (M) 5.4-6.3 x 3.6-4.5 5.85
Hydnellum complicatum (B) 4-5 x 3-5 4.50
Hydnellum concrescens (M) 5.4-6.1 x (3.6-)4-4.5 dep
Hydnellum cristatum (B) 5-6 x 4-5 5.50
Hydnellum cruentum K.A.Harrison (B) 4-5 x 3-4 4.50
Hydnellum cumulatum (M) 4.3-5.6 x 3.6-4.3 4,95
Hydnellum diabolus (B) 6-7 x 5-6 6.50
Hydnellum earlianum (B) 5=6:% 425 5.50
Hydnellum fennicum (M) 6.3-7.6 x 4.5-5.2 6.95
Hydnellum ferrugineum (M) (5.4-)5.8-6.3 x 3.6-4.5 6.05
Hydnellum ferrugipes (B) 5-7 x 5-6 6.00
Hydnellum fuligineoviolaceum (M) 5.4-6.5 x 4-4.7(-5.4) 5:95
Hydnellum geogenium (M) 4,5-5.2 x 3.1-3.6 4.85
Hydnellum glaucopus (M) (5-)5.4-5.8(-6.3) x (3.6-)4-4.5 5.60
Hydnellum gracilipes (M) 4.3-4.6 x 2.7-3.6 4.45
Hydnellum joeides (M) 5.4-5.8 x 3.6-4.2 5.60
Hydnellum lepidum (M) 5.8-6.3 x 3.6-4.3 6.05
Hydnellum lundellii (M) 4.9-5.8 x 3.6-4.2 5.35
Hydnellum martioflavum (M) 5-6.3 x 3.6-4.5 5.65
Hydnellum peckii (M) 4.9-5.4 x 3.8-4 5215
Hydnellum pineticola (B) 5-7 x 4-6 6.00
Hydnellum piperatum (B) 4-6 x 4-5 5.00
Hydnellum scabrosum (M) (5.4-)6.3-7.3 x (3.6-)4-5 6.80
Hydnellum scleropodium (B) 4-6 x 3-4 5.00
Hydnellum spongiosipes (B) 6-7 x 5-6 6.50
Hydnellum suaveolens (M) 4-5 x 3-3.6 4.50
Hydnellum subsuccosum (B) 5-6 x 4-6 5.50
Hydnellum versipelle (M) 4.5-5.5 x 3.5-4.5 5.00
Hydnellum underwoodii (B) 5-7 x 5-6 6.00
Sarcodon atroviridis (B) 8-9 x 7-8 8.50
Sarcodon excentricus R.E.Baird (B) 8-9 x 6-8 8.50
Sarcodon harrisonii R.E.Baird (B) 7-9 x 6-8 8.00
Sarcodon leucopus (M) (6.7-)7.2-7.6(-9) x 4.5-5.6 7.40
Sarcodon imbricatus (M) 7.2-8.2 x 4.9-5.4 7.70
Sarcodon scabripes (B) 8-10 x 7-9 9.00
Sarcodon squamosus (J) 7.2-8.2 x 4.9-5.4 7.70

marker. Our general experience with the ITS region for thelephoralean target genera
is that species are extremely well separated and the internal variation surprisingly low,
even when a large number of specimens from both Europe and America are considered.
On the other hand, the genetical difference among species is moderate to high, making
alignments difficult and prone to ambiguities. In our ITS analyses we chose to remove
ambiguous regions, thus halving the number of nucleotide positions suggested by

Reassessment of the generic limits for Hydnellum and Sarcodon 45

automatic alignment through MAFFT. This seems to have affected the ML analyses
most. However, the ITS analyses only served to position neotropical Sarcodon species
and the results clearly show that they belong to a separate lineage.

Otto (1997) suggested that Aydnum auratile is a later synonym of
Hydnum aurantiacum and that the species we now call Hydnellum aurantiacum should
be named Hydnellum floriforme (Schaeff.) Banker. The name change is based on a
reinterpretation of Batsch’s original illustration, which, according to Otto, clearly shows
the same species as Hydnum auratile. In phylogenetic analyses H. aurantiacum and
H. auratile are sister taxa and during our study we have sequenced several specimens
identified as H. auratile that turned out to be H. aurantiacum. Thus separating these
species can be hazardous and to interpret illustrations must be even harder. We currently
do not accept this unfortunate name change.

The present study will serve as the basis for further exploration of species limits
within Hydnellum and Sarcodon. As has been demonstrated for the genera, many
species interpretations are in need of revision. Over the years we have found numerous
specimen misidentifications as well as specimens that could not be assigned to pre-
existing names. A closer inspection of the ITS tree in Fig. 3, where we let the terminals
retain the identifications given in GenBank, shows some examples. The American
sequence of Sarcodon joeides (KC571772) does not cluster with the European
representative of the same species (MK602751) and the American sequence named
Hydnellum earlianum seems to be identical to what is in Europe called H. auratile.
Considering that many stipitate hydnoid species are red-listed and used as indicators
of forests in need of conservation (Ainsworth 2005, Nitare 2019), it is of utmost
importance to sort out the taxonomy of these species.

Acknowledgements

This study was supported by grants from ArtsDatabanken, Norway, to KH Larsson
(ADB54-09), from Artdatabanken, Sweden, to E Larsson (2014-152 4.3), and from
Estonian Research Council to U Koljalg (IUT20-30). We also acknowledge support
to S Svantesson from Kungliga Vetenskaps- och Vitterhetssamhallet i Goteborg and
from Kapten Carl Stenholms donatationsfond. We are grateful to many dedicated
mycologists in Norway, Sweden and Finland for sending valuable collections. We are
especially grateful to Johan Nitare for sharing with us his outstanding knowledge of
stipitate hydnoid fungi and for duplicates from his herbarium. We also thank Martyn
Ainsworth and Terry Henkel whose thorough reviews improved this paper considerably.

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